A painting of Britain's King Richard III by an unknown artist is displayed in the National Portrait Gallery in London. / Leon Neal, AFP/Getty

by Naomi Westland, Special for USA TODAY

by Naomi Westland, Special for USA TODAY

LEICESTER, England -- It took five centuries but the mystery over what happened to the remains of England's last Plantagenet king, Richard III, is finally solved after scientists at the University of Leicester confirmed a skeleton buried under a city parking lot is the much maligned king vilified by Shakespeare.

Monday's announcement brings to a close the long-debated question of what followed the killing of Richard III on the battlefield 528 years ago. Further tests on the skeleton that suggest that accounts of his withered arm are inaccurate may help to restore his reputation, historians say.

"This is a historic moment and the history books will be rewritten," said Philippa Langley, originator of the search and a member of the Richard III Society. "We have searched for Richard and found him ‚?? now it is now time to honor him."

The remains will be reburied in Leicester Cathedral.

The skeleton was found in September in a University of Leicester dig on the site of a medieval church, Grey Friars, now a parking lot for a social services office. It had what was thought to be a metal arrow in its back ‚?? in fact, more likely to be a Roman nail -- and head injuries. Both are consistent with records of how Richard III was killed at the Battle of Bosworth in 1485. It also had a curved spine, fitting with reports that one of Richard III's shoulders was higher than the other.

Over the past four months, the remains have been subjected to a series of tests, including a computed‚??tomography (CT) scan and radio carbon dating. Specialists in medieval battles and weaponry advised on the kinds of instruments that may have caused the damage to the skull.

Scientists carried out tests on DNA from the remains, comparing it with a sample from Canadian furniture-maker Michael Ibsen, 55, now confirmed along with his siblings as the 17th-generation descendant of Richard III's sister, Anne of York.

Ibsen said he was stunned when he heard the news yesterday. "I'm still trying to digest it. I've been aware of the on-going tests but I never thought there would be an almost perfect match with the DNA," he said.

A number of other men identified as descendants of Richard's great-great-grandfather, Edward III, through his son John of Gaunt ‚?? who both would share the same Y chromosome as Richard III -- also donated DNA to the project to confirm the male line.

The analysis of the male line is still in its early stages but Turi King, a University of Leicester geneticist, hopes it will provide a complete picture on both lines of descent.

"The DNA sequence obtained from the Grey Friars skeletal remains was compared with the two maternal line relatives of Richard III ‚?? we were very excited to find that there is a DNA match between the maternal DNA from the family of Richard III and the skeletal remains at the Grey Friars dig," King said.

Skeletal analysis carried out by Jo Appleby, an osteoarcheologist at the university, showed that the individual was male and in his late 20s to late 30s. Richard III was 32 when he was killed. Trauma to the skeleton shows cause of death was the result of one or two wounds to the back of the skull that could have been made by a sword or halberd, a type of ax. This is consistent with accounts of Richard being killed after receiving a blow to the back of his head. The skeleton also showed a number of non-fatal injuries ‚?? to the head, rib and one to the pelvis believed to be from a wound through the right buttock -- which Appleby says may have been inflicted as humiliation after death.

"The skeleton has a number of unusual features: its slender build, the scoliosis and the battle-related trauma," she said. "All these are highly consistent with the information that we have about Richard III in life and about the circumstances of his death. Taken as a whole, the skeletal evidence provides a highly convincing case for identification as Richard III."

Richard III reigned for two years until his death at Bosworth, the last major battle of the dynastic struggle known as the Wars of the Roses fought on and off from 1455. The last English king to be killed in battle, Richard's death brought about the end of three centuries of Plantagenet rule and ushered in the Tudors -- starting with Henry VII and including Henry the VIII -- who reigned for the next 117 years.

Although Richard remains the prime suspect in the disappearance of his nephews -- the princes in the tower who vanished after their uncle assumed the throne -- many academics and historians hope the positive identification will go some way to restoring Richard's reputation.

Modern views of the medieval king have been heavily influenced by William Shakespeare's portrayal of him in the play Richard III, historians say.

Shakespeare shows Richard as a power-hungry, Machiavellian scoundrel who goes around murdering anyone who stands in the way of his ascent to power. He depicts him as having a withered arm but the new scientific evidence discredits this description as both of the skeleton's arms are the same length.

Langley said Richard III was a progressive leader who pioneered a system of bail for those arrested, the legal principles of presumption of innocence until proven guilty and blind justice, and that he introduced books to England.

"It's ironic that he is guilty until proven innocent of the disappearance of the princes even though there is no evidence against him," she said.

She said the "winds of change are blowing" on his reputation and pledged to seek out the truth.

"The two-dimensional caricature promoted by the Tudors will be no more," she said.

Ibsen hopes the identification of his royal predecessor will stimulate people to take a wider interest in Richard III, and not just of his short reign. "I hope it will lead to a broader evaluation of his life as a whole," he said.

Jack Douglass, 19, a history student at the University of Leicester, said he was thrilled by the discovery

"It's all incredibly exciting," he said. "People tend to think of Richard III in black and white but one of the best things is that it will open up the debate around him and now people in the real world will start to talk about him, not just academics."

At the entrance to the unassuming parking lot below which the skeleton was discovered, passersby stop and crane their necks to get a better view of the trench that was Richard's resting place until four months ago.

Anthony McBride, 64, standing with his hands shoved into his pockets against the cold, was pleased to know that the remains were the king's.

"Nottingham has Robin Hood but he's just a myth," he said. "We have the real thing. A real king who died bravely in battle, let down by the people he trusted. It would have been so different if he'd won, that horrible Henry VIII wouldn't have come to power."